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Osteology, Natural History Notes, and Phylogenetic Relationships of the Poorly Known Caribbean Frog Leptodactylus Nesiotus (Anura, Leptodactylidae) María Laura Ponssa

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Osteology, Natural History Notes, and Phylogenetic Relationships of the Poorly Known Caribbean Frog Leptodactylus Nesiotus (Anura, Leptodactylidae) María Laura Ponssa University of Richmond UR Scholarship Repository Biology Faculty Publications Biology 10-12-2010 Osteology, Natural History Notes, and Phylogenetic Relationships of the Poorly Known Caribbean Frog Leptodactylus nesiotus (Anura, Leptodactylidae) María Laura Ponssa Michael J. Jowers Rafael O. de Sá University of Richmond, [email protected] Follow this and additional works at: http://scholarship.richmond.edu/biology-faculty-publications Part of the Biology Commons, Population Biology Commons, Terrestrial and Aquatic Ecology Commons, and the Zoology Commons Recommended Citation Ponssa, María Laura, Michael J. Jowers, and Rafael O. de Sá. "Osteology, Natural History Notes, and Phylogenetic Relationships of the Poorly Known Caribbean Frog Leptodactylus nesiotus (Anura, Leptodactylidae)." Zootaxa 2646 (October 14, 2010): 1-25. This Article is brought to you for free and open access by the Biology at UR Scholarship Repository. It has been accepted for inclusion in Biology Faculty Publications by an authorized administrator of UR Scholarship Repository. For more information, please contact [email protected]. Zootaxa 2646: 1–25 (2010) ISSN 1175-5326 (print edition) www.mapress.com/zootaxa/ Article ZOOTAXA Copyright © 2010 · Magnolia Press ISSN 1175-5334 (online edition) Osteology, natural history notes, and phylogenetic relationships of the poorly known Caribbean frog Leptodactylus nesiotus (Anura, Leptodactylidae) MARÍA LAURA PONSSA1, MICHAEL J. JOWERS2, & RAFAEL O. DE SÁ3 1CONICET. Fundación Miguel Lillo, San Miguel de Tucumán, Tucumán, Argentina. E-mail: [email protected] 2Division of Ecology and Evolutionary Biology, Institute of Biomedical and Life Sciences, University of Glasgow, Glasgow, UK. E-mail: [email protected] 3Department of Biology, University of Richmond, Richmond, Virginia, 23173, US. E-mail: [email protected] Abstract The Leptodactylus melanonotus group consists of 15 species, but references to skeletal characters are available for only three species: L. leptodactyloides, L. melanonotus, and L. diedrus. Leptodactylus nesiotus is a member of the melanonotus group known only from the type locality, Bonasse swamp, on the Southwestern peninsula of Trinidad, Trinidad and Tobago. This species has been categorized as vulnerable given its restricted distribution. Herein, we report the adult osteology of L. nesiotus, the skeletal characters are compared with the available data from other Leptodactylus species. A phylogenetic analysis recovers a paraphyletic L. melanonotus group relative to the L. latrans group. A monophyletic “latrans-melanonotus” clade is supported by five synapomorphies. L. nesiotus is recovered as the sister species of L. validus, a relationship supported by two synapomorphies: T-shaped terminal phalanges and a dark-colored stripe on the outer surface of arm. In addition, we report on the ecology of this poorly known species. Key words: skeleton, ecology, phylogenetic relationships, Leptodactylus nesiotus Resumen El grupo L. melanonotus del género Leptodactylus incluye 15 especies, aunque sólo existen reportes de caracteres osteológicos para tres de ellas: L. leptodactyloides, L. melanonotus y L. diedrus. Leptodactylus nesiotus es un miembro del grupo L. melanonotus que ha sido categorizado como vulnerable debido a su distribución restringida, ya que es conocido sólo de la localidad tipo: La ciénaga de Bonasse, sudoeste de la península de la isla Trinidad, Trinidad y Tobago. En este trabajo se aportan datos sobre la osteología de adultos de L. nesiotus, los cuales son comparados con lo observado en otras especies del género, y se sumaron a una matriz previamente elaborada. Esta fuente de caracteres se utilizó para realizar un análisis de parsimonia y proponer una hipótesis filogenética. El grupo L. melanonotus resultó parafilético respecto al grupo L. latrans. El clado latrans-melanonotus es apoyado por cinco sinapomorfías. Se infiere a Leptodactylus nesiotus como la especie hermana de L. validus, relación apoyada por dos sinapomorfías: falanges terminales en forma de T, y presencia de banda oscura en el borde externo del brazo. Además se reportan observaciones sobre la ecología de esta especie pobremente conocida. Introduction The genus Leptodactylus (Fitzinger 1826) consists of 87 species (Frost 2009) that have been traditionally clustered into five species groups Leptodactylus fuscus, Leptodactylus melanonotus, Leptodactylus latrans, Leptodactylus pentadactylus, and Leptodactylus marmoratus (Heyer 1969a). Phylogenetic studies of the genus have been limited, only including a few species of each group (Maxson & Heyer 1988; Heyer 1998; Larson & de Sá 1998; de Sá, Heyer & Camargo 2005; Heyer et al. 2005) or focusing on a single group (Ponssa 2008). Leptodactylus nesiotus (Heyer 1994) is currently considered a member of the L. melanonotus group. This species has been categorized as vulnerable given its restricted distribution, known only from the Accepted by M. Vences: 9 Sep. 2010; published: 14 Oct. 2010 1 type locality, in the Bonasse Swamp on the southwestern peninsula of Trinidad, Trinidad and Tobago (IUCN 2006). Osteology has been traditionally and widely used in amphibian phylogenetic studies. References to skeletal features and osteological characters of Leptodactylus are available for: L. chaquensis (Heyer 1998; Perotti 2001), L. insularum (Heyer 1998), L. laticeps (Ponssa 2006), L. pentadactylus (Heyer 1969b; Heyer 1998), L. riveroi (Heyer 1998), L. lauramiriamae (Heyer & Crombie 2005), L. silvanimbus (Heyer 1998), species of the L. fuscus group (Heyer 1998; Ponssa & Lavilla 1998; Ponssa 2008), species of the subgenus Lithodytes (Heyer 1974; 1998; Ponssa & Heyer 2007), and for the Leptodactylus genus in general (Lynch 1971). Among the 15 species in the L. melanonotus group, osteological characters have been previously reported only for L. leptodactyloides, L. melanonotus, and L. diedrus (Heyer 1998). Osteological characters within this group are useful for species diagnoses, to understand patterns of morphological evolution, and can contribute to assess phylogenetic relationships. Herein, we report on the adult osteology of L. nesiotus and compare it with other available data for the genus. A phylogenetic analysis was performed to assess the relationships of this species to other Leptodactylus species and to test the monophyly of the traditional L. melanonotus species group. Furthermore, we report observations on the ecology and natural history of this poorly known species. Material and methods Three Leptodactylus nesiotus males and one female (N = 4) were collected at Bonasse swamp (10°05’41.55”N, 61°49’53.35”W; 10 m.s.n.m.), Trinidad (Trinidad and Tobago), on July 3, 2004, by M. J. Jowers and R. Campbell-Palmer. The specimens were taken to the laboratory for further observations. They were housed in tanks (0.95 cm x 35 cm x 35 cm) with a central mud islet with plenty of shrub branches, wood, few stones, leaf litter, and water to best resemble their natural habitat. The swamp is easily accessed through the Southern Main road that divides the swamp into two separate areas. To the Northern side of the road, the swamp reaches its maximum water capacity, about a meter deep. Here, the swamp (about half a hectare) is composed of few tall trees growing in the water and smaller trees on islet formations found around the back and centre of the swamp. The swamp is surrounded by tall reeves growing at the side of the road on the Northern section. The Southern side of the swamp is more open and resembles a field of tall grasses after a heavy rainfall. The water rarely reaches more than a foot in depth and the area was occasionally dry. Frogs were not heard calling and thus surveys were not carried out in this area. For the morphological analysis two Leptodactylus nesiotus specimens, a female (USNM 558322) and a male (USNM 558321) were cleared and double stained following Wassersug’s (1976) protocol. Skull measurements were taken with Image Tool software (Fig. 1). The terminology for cranial and postcranial osteology follows Trueb (1973; 1993) and Trueb et al. (2000). Terminology of digits and carpal osteology follows Fabrezi (1992), olfactory region follows that of Pugener and Maglia (2007) and Maglia et al. (2007), laryngeal morphology follows Trewavas (1933), and sesamoids terminology follows Ponssa et al. (2010). Fingers are numbered II–V following Fabrezi and Alberch (1996). Osteological data of species of the L. fuscus group follows Ponssa (2008) whereas data for L. colombiensis, L. melanonotus, L. validus, and L. silvanimbus (species in the L. melanonotus species group) follow Heyer (1998) and Ponssa (2008). The osteology of L. laticeps was included as a representative species of the L. pentadactylus group (Ponssa 2006). Herein, the L. latrans group refers to the traditionally known L. ocellatus group, following Lavilla et al. (2010). Characters that resulted from the description were coded and combined with those previously published for the Leptodactylus fuscus group (Ponssa 2008) (Appendix 1). A maximum parsimony (MP) analysis (i.e., traditional search, with 2000 different addition sequences to the tree bisection-reconnection branch swapping method (TBR), retaining 100 trees per replication; internal branches were considered unsupported and collapsed during searches if any possible states were shared between ancestor and descendent nodes -min. length = 0 option-) was performed in TNT (Goloboff et al. 2 · Zootaxa 2646 © 2010 Magnolia Press PONSSA ET AL. 2003a).
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